ULTRASTRUCTURAL STUDY ON SPORE WALL MORPHOGENESIS IN LYCOPODIUM CLAVATUM (LYCOPODIACEAE)

Spore wall morphogenesis of Lycopodium clavatum was observed by transmission electron microscopy. The spore plasma membrane indicates the reticulate spore sculpture shortly after meiosis. The mature spore wall of this species consists of two layers, inner endospore and outer exospore. There is no perispore in the sporoderm of this species. The exospore formation begins during the tetrad stage; and this layer is divided into two distinct sublayers, an outer lamellar layer and an inner granular layer. The lamellar layer is formed on the sculptured spore plasma membrane. Additional lamellae attach to this layer in a centripetal direction. For that reason, this layer may be derived from spore cytoplasm. The granular layer is formed only in the proximal region following lamellar layer formation, and it also may be derived from spore cytoplasm. The endospore is formed lastly and seems to be derived from spore cytoplasm as well. Accordingly, the spore sculpture of this species may be under the genetic control of the spore nucleus.     

ULTRASTRUCTURAL STUDY ON MICROSPORE WALL MORPHOGENESIS IN ISOETES JAPONICA (ISOETACEAE)

Spore wall morphogenesis of the microspore of Isoetes japonica was studied by transmission electron microscopy. The microspore wall consists of four layers: the perispore, outer exospore, inner exospore, and endospore. The perispore consists of electron-dense materials. The exospore is divided into outer and inner sections, with a large gap between the two. The outer exospore appears as an undulating plate consisting of tripartite lamellae with homogeneous sporopollenin. The inner exospore consists of an accumulation of tripartite lamellae on the microspore cell membrane. Immediately after meiosis, the tripartite lamellae of the outer exospore forms around the microspore. The lamellated inner exospore forms next, which adheres to the cell membrane of the microspore. The deposition of homogeneous sporopollenin material on the tripartite lamellae causes the plates of the outer exospore to thicken. Some homogeneous material may also be deposited on the inner exospore. Lastly, the electron-dense perispore is deposited on the outer exospore, and the electron-lucent endospore forms beneath the inner exospore. We conclude that the lamellae of the outer exospore, inner exospore, and endospore are formed and derived, in that order, from the gametophytic microspore cytoplasm. The homogeneous sporopollenin material of the outer exospore and perispore may be derived from the sporophytic tapetal cytoplasm.     

朝鲜介蕨孢子周壁发育的研究

利用光镜、扫描电镜和透射电镜对朝鲜介蕨[Dryoathyrium coreanum(Christ)Tagawa=Lunathyrium coreanum(Christ)Ching]孢子周壁的发育规律进行了研究。结果表明,朝鲜介蕨孢子两侧对称,单裂缝,表面具粗大的脊状褶皱,褶皱形成网状或拟网状纹饰。孢壁包括内壁、外壁和周壁。孢子外壁表面光滑,在四分孢子时期就已发育成熟。四分孢子分离后,周壁开始形成,周壁来源于孢子囊的绒毡层,是由原质型绒毡层的残余物在外壁上沉积而成。成熟的周壁很厚,可分为外层和内层。周壁内有大的空腔,主要是由周壁外层向外隆起形成的,隆起进而形成了孢子的脊状褶皱和表面纹饰。     

瓦韦孢子壁的结构和发育的研究

利用光镜、扫描电镜和透射电镜对水龙骨科（Polypodiaceae）瓦韦(Lepisorus thunbergianus (Kaulf.) Ching)孢子壁的结构和发育进行了研究。研究结果表明瓦韦孢子两侧对称、单裂缝,表面具波纹状纹饰。孢壁从内到外由内壁、外壁和周壁三部分构成。外壁来源于绒毡层物质,由外壁内层和外壁外层构成,外壁外层表面的波纹状纹饰形成孢子表面的纹饰轮廓。周壁薄,紧贴外壁表面,由2层片状结构叠合而成。在外壁外层形成过程中,孢子表面和周围出现较多小球。本文探讨了孢壁各层的结构、来源和发育过程,为蕨类植物系统学和孢粉学研究积累资料。     

乌蕨孢子壁的形成和发育

利用光镜、扫描电镜和透射电镜对鳞始蕨科（Lindsaeaceae）乌蕨（Stenoloma chusanum Ching）孢壁的形成和发育进行了研究。结果表明乌蕨孢子两侧对称、单裂缝,表面具疣状纹饰。孢壁由内壁、外壁和周壁三部分构成。外壁在四分体阶段已基本形成,其表面光滑,质地均匀,由孢粉素形成。周壁是由绒毡层残余物在外壁表面沉积形成,可分为周壁内层、周壁中层和周壁外层三部分。在周壁中层与外层之间有一层均匀的空间。最后,本文探讨了孢壁的形成和发育规律,研究结果对揭示孢子纹饰和孢壁各层的形成过程、来源和稳定性有重要的意义,并为孢粉学和系统学研究提供基础资料。     

乌蕨孢子壁的形成和发育

利用光镜、扫描电镜和透射电镜对鳞始蕨科(Lindsaeaceae) 乌蕨( Stenoloma chusanum Ching) 孢壁的形成和发育进行了研究。结果表明乌蕨孢子两侧对称、单裂缝, 表面具疣状纹饰。孢壁由内壁、外壁和周壁三部分构成。外壁在四分体阶段已基本形成, 其表面光滑, 质地均匀, 由孢粉素形成。周壁是由绒毡层残余物在外壁表面沉积形成, 可分为周壁内层、周壁中层和周壁外层三部分。在周壁中层与外层之间有一层均匀的空间。最后, 本文探讨了孢壁的形成和发育规律, 研究结果对揭示孢子纹饰和孢壁各层的形成过程、来源和稳定性有重要的意义, 并为孢粉学和系统学研究提供基础资料。     

The tapetum inSchizaea pectinata (Schizaeaceae) and a comparison with the tapetum inPsilotum nudum (Psilotaceae)

A combination tapetum consisting of a cellular, parietal component and a plasmodial component occurs inSchizaea pectinata. A single, tapetal initial layer divides to form an outer parietal layer which maintains its cellular integrity until late in spore wall development. The inner tapetal layer differentiates into a plasmodium which disappears after the outer exospore has developed. In the final stages of spore wall development, granular material occurs in large masses and is dispersed as small granules throughout the sporangial loculus. No tapetal membrane develops. Comparisons are drawn with the combination tapetum found inPsilotum nudum.     

紫萁孢子囊发育中多糖和脂滴的细胞化学观察

采用光镜、透射电镜和细胞化学技术,对紫萁孢子囊发育过程中孢壁的超微结构和孢子囊内多糖和脂滴的分布及其动态变化进行研究,以探讨紫萁孢子囊发育过程中多糖和脂滴的代谢特征,为蕨类孢子发生的研究提供基础资料。结果表明:(1)紫萁孢子囊由1层囊壁细胞、2层绒毡层和产孢组织构成。(2)紫萁孢子壁由发达而分2层的外壁(外壁内层和外壁外层)和薄的不连续的周壁构成,由外壁形成棒状纹饰的轮廓;孢子外壁内层由多糖类物质构成,外壁外层和周壁均含有脂类物质。(3)在紫萁孢原细胞中观察到少量脂滴;随着紫萁孢壁的形成,囊壁细胞中淀粉粒的大小逐渐变小、数目先增加后减少,它们转运到内层绒毡层原生质团并转化为孢粉素前体物质,再穿过原生质团内膜表面进入囊腔,成为孢粉素团块或以小球形式填加到孢子表面形成孢壁。(4)紫萁孢子囊将多糖类营养物质转化为脂类,以脂滴的形式储藏在孢子中。     

红盖鳞毛蕨孢子发育的超微结构和细胞化学研究

采用透射电镜和细胞化学技术对红盖鳞毛蕨(Dryopteris erythrosora(Eaton)O.Ktze.)的孢子发育过程进行了研究,根据超微结构和细胞化学特征可将其孢子发育过程分为3个阶段:(1)孢子母细胞及其减数分裂阶段:孢子母细胞壳在孢原细胞末期开始形成,位于孢子母细胞及其减数分裂形成的四分体外侧,PAS反应显示其为多糖性质,与胼胝质壁为同功结构;在减数分裂形成的四分孢子之间产生孢子外壳,从功能、形成位置和时间上看与胼胝质壁相似,但苏丹黑B反应显示其可能含有脂类物质,与孢子母细胞壳和胼胝质壁不同。(2)孢子外壁形成阶段:外壁为乌毛蕨型(Blechnoidal-type),由薄的多糖性质的外壁内层和表面平滑的孢粉素外壁外层构成;小球参与外壁外层的形成,组织化学分析显示小球的中央区域和外壁外层内侧部分由红色(多糖)变为黄色,小球的表面区域和外壁外层部分始终被染成黑色(脂类),可知小球与外壁同步发育。(3)孢子周壁形成阶段:周壁为凹陷型(Cavate-type),包括2层,内层薄,紧贴外壁,外层隆起形成孢子脊状褶皱纹饰的轮廓,以少见的向心方向发育;苏丹黑B和PAS反应观察周壁被染成橙色,推测其可能由多糖等成分构成;孢子囊壁细胞参与周壁的形成。本研究为揭示蕨类植物孢子发生的细胞学机制提供了新资料。     

团扇蕨孢子发生和发育的显微观察

利用光学显微镜对膜蕨科（Hymenophyllaceae）团扇蕨(Gonocormus minutus(Blume) Bosch)孢子的发生和发育进行了观察。研究结果表明：团扇蕨孢子为多边圆形,三裂缝不明显,外壁表面光滑,周壁薄,紧贴外壁表面,由周壁形成乳头状或颗粒状纹饰。在外壁形成后期,孢子表面和囊腔中出现大量小球;在周壁形成时期,孢子表面和周围出现较多小球体;小球和小球体参与孢子壁的形成。团扇蕨绒毡层为混合型,内层为周原质团绒毡层;外层为腺质型绒毡层。本文为膜蕨科系统演化和发育生物学研究提供依据。     

Spore morphology and ultrastructure of Cyathea (Cyatheaceae,Pteridophyta) species from southern South America

Spore sculpture and wall structure of eight Cyathea (Cyatheaceae) species from southern South America were studied using light microscopy (LM), scanning electron microscopy (SEM) and transmission electron microscopy (TEM) techniques. Two layers, i.e. an inner and an outer layer, were observed in the perispore. The inner layer has two strata: the inner stratum is attached to the exospore and composed of rodlets tangentially oriented to the spore surface and randomly intermixed; the outer stratum consists of a three-dimensional network of rodlets with either free or fused distal edges forming spinules. The outer layer is thin, darkly contrasted and covers the rodlets. In most cases, the exospore has two layers and a pitted surface. In Cyathea atrovirens, the exospore surface is smooth, while in C. delgadii and C. myriotricha it is verrucate. The homogeneity of perispore features within the genus Cyathea is evident, while exospore features are heterogeneous. The exospore has different kinds of surface-structures that are of potential interest for assessing evolutionary trends within the group.     

Microsporidian Spore Wall: Ultrastructural Findings on Encephalitozoon hellem Exospore

A study of the spore wall of Encephalitozoon hellem was performed on thin sections, freeze-fracture, and deep-etched samples to obtain information on spore wall organization and composition. Our observations demonstrate that the spore wall is formed by an inner 30–35 nm electron-lucent endospore and an outer 25–30 nm electron-dense exospore. The exospore is a complex of three layers: an outer spiny layer, an electron-lucent intermediate lamina and an inner fibrous layer. Freeze-fracture and deep-etching techniques reveal that the intermediate lamina and the inner fibrous layer result from the different spatial disposition of the same 4-nm thick fibrils. In thin sections the endospore reveals a scattered electron-dense material that appears in the form of trabecular structures when analyzed in deep-etched samples. The presence of chitin in the exospore is discussed.     

海金沙孢子壁结构和发育的研究

利用光镜、扫描电镜和透射电镜对海金沙科(Lygodiaceae)海金沙[Lygodium japonicum (Thunb.) Sw.]孢壁的形成和发育进行了研究.结果表明:海金沙孢子壁由内壁、外壁和周壁3部分构成.外壁由2层构成,即薄的内层和厚的外层,其中外层是在四分体分离前通过孢粉素的逐层沉积并浓缩凝聚而形成的均质层,其表面具不明显的疣状突起.周壁由绒毡层残余物在外壁表面逐层沉积形成,可分为周壁内层、周壁中层和周壁外层3部分;周壁中层具辐射状排列的长条形成分,周壁外层形成瘤状纹饰的轮廓.本研究为孢粉学和蕨类植物系统演化分析提供基础资料.     

蕨类植物紫萁绒毡层细胞凋亡的细胞学特征

绒毡层凋亡过程是小孢子发生中的重要事件,以往的研究主要集中在被子植物,蕨类植物尚未见此方面的报道。该研究首次采用透射电镜和免疫荧光技术对蕨类植物紫萁(Osmunda japonica Thunb.)绒毡层细胞凋亡的细胞学过程进行了观察,以明确紫萁绒毡层细胞的发育类型和凋亡特征,为蕨类植物绒毡层细胞凋亡的深入研究以及孢子发育研究提供依据。结果显示：(1)紫萁的绒毡层属于复合型,即外层绒毡层为分泌型,该层细胞发育过程中液泡化,营养物质被吸收;内层绒毡层为原生质团型,经历了细胞凋亡的过程。(2)绒毡层内层细胞在凋亡过程中细胞壁和细胞膜降解,细胞质浓缩且空泡化;细胞核内陷、变形,染色质浓缩凝聚,形成多数小核仁,DAPI荧光由强变弱;线粒体、质体、内质网、高尔基体等细胞器逐渐退化,液泡中多包含纤维状物、絮状物、黑色嗜锇颗粒和小囊泡等;出现多泡体、多膜体和细胞质凋亡小体,上述特征与种子植物绒毡层凋亡特征基本一致。(3)与种子植物相比,紫萁绒毡层的细胞凋亡开始得早,在整个凋亡过程中没有核凋亡小体的产生;除了产生孢粉素外,绒毡层细胞内产生了大量的丝状物质、絮状物质和电子染色暗的颗粒物,这些物质可能用于...     

蜈蚣草孢壁发育及其系统学意义

利用光镜、扫描电镜和透射电镜对凤尾蕨科(Pteridaceae)蜈蚣草(Pteris vittata L.)孢壁的形成和发育进行研究。结果表明:蜈蚣草孢子四面体型,极面观钝三角圆形,赤道面观半圆形或超半圆形,近极面具瘤状纹饰和近极脊,远极面具脊并连成网状,具赤道环;孢子具乌毛蕨型外壁,由外壁外层构成纹饰的轮廓;实心型周壁由2层构成,且内层薄、外层具小球体。结合孢子外壁和周壁的发育特征,认为凤尾蕨科与裸子蕨科和水蕨科的亲缘关系较近,支持将裸子蕨科和水蕨科置于凤尾蕨科。     

狗脊孢壁发育超微结构的研究

鳞毛蕨型孢子类型众多,初步研究表明形态相似的孢子类型其孢壁发育特征存在差异,因此有必要对各代表类群的孢壁发育进行深入地研究。该文利用透射电镜对乌毛蕨科(Blechnaceae)狗脊(Woodwardia japonica)孢壁结构和发育的超微结构进行研究。结果表明:(1)狗脊孢子囊的结构由外向内分别为孢子囊壁细胞、两层绒毡层细胞和孢子母细胞;(2)狗脊孢子具乌毛蕨型(Blechnoid type)外壁,表面光滑,由两层构成,裂缝区域具辐射状的槽;(3)周壁属于空心型(cavity type),由四层构成,从内向外分别为P1、P2、P3和P4层,前三层叠合在一起,层间有不同程度的空隙,P4层与前三层之间具有明显而连续的空腔,并隆起形成片状褶皱纹饰;(4)有小球体和小杆共同参与孢子周壁的形成,周壁部分或全部来源于孢子囊壁细胞。综上所述,狗脊孢子与同属于鳞毛蕨型的贯众(Cyrtomium fortunei)和朝鲜介蕨(Dryoathyrium coreanum)孢壁的发育在周壁结构、周壁各层的发育顺序、周壁来源和参与成壁的特征物质等方面存在差异。该研究有利于进一步理解蕨类植物孢壁所蕴含的分类和演化上的科学意义和价值。     

Ultrastructure of developing basidiospores in <Emphasis Type="Italic">Rhizopogon roseolus</Emphasis> (= <Emphasis Type="Italic">R. rubescens</Emphasis>)

The ultrastructure of developing basidiospores in Rhizopogon roseolus is described. When viewed in the fruiting body chamber using scanning electron microscopy, basidiospores appear narrowly ellipsoid and have smooth walls. Eight basidiospores are usually produced on the apex of each sterigma on the basidium. Transmission electron micrographs showed that basidiospores formed by movement of cytoplasm (including the nuclei) via the sterigmata, and then each basidiospore eventually became separated from its sterigma by an electron-lucent septum. The sterigma and basidium subsequently collapsed, resulting in spore release. Freshly released spores retained the sterigmal appendage connected to the collapsed basidium. After spore release, the major ultrastructural changes in the spore concerned the lipid bodies and the spore wall. During maturation, lipid bodies formed and then expanded. Before release, the spore wall was homogeneous and electronlucent, but after release the spore wall comprised two distinct layers with electron-dense depositions at the inner wall, and the dense depositions formed an electron-dense third layer. The mature spore wall complex comprised at least four distinct layers: the outer electron-lucent thin double layers, the mottled electron-dense third layer, and the electron-lucent fourth layer in which electron-lucent granular substances were dispersed.     

Spore morphology and wall ultrastructure of Hymenophyllaceae Link (Pteridophyta) from north-west Argentina

The family Hymenophyllaceae is represented in the study area by six species in two genera, Hymenophyllum J. E. Smith and Trichomanes L. The study was based on herbarium material and spores were studied under light microscope (LM), scanning electron microscope (SEM) and transmission electron microscope (TEM). Both genera have trilete spores, 23 to 45 μm in equatorial diameter, with an ornamentation of echinulae and cones in Hymenophyllum and of verrucae, gemmae and granules in Trichomanes. Mature spores have a sporoderm composed of a perispore, an exospore and a fibrillar endospore; the exospore is 0.5 to 2.5 μm thick, compact and with an irregular margin. In some cases radial channels and other channels associated with the middle and inner parts of the laesurae were evident. A series of cavities filled with an opaque content line the inner margin of the exospore. The perispore is 20 to 400 nm thick and unevenly differentiated along the surface of a same spore. Under TEM, two main differentially contrasted portions could be distinguished: a dark massive portion with structural components could not be distinguished, and a light portion with several plates arranged in piles. The inner surface of the perispore exhibit short scales. Globules are immersed within the perispore at some depth from the perispore surface and others connected to it by structural threads. The spore characters observed including shape, ornamentation, laesurae length and wall structure are useful in distinguishing the two genera studied, but less useful in differentiation at the species level.     

Crepidula beklemishevi gen. et sp. n. and Dimeiospora palustris gen. et sp. n. (Microspora: Amblyosporidae)--new microsporidian genera and species from blood-sucking mosquitoes (Diptera: Culicidae) from the south of the western Siberia

Microsporidia parasitizing the adipose body of mosquito larvae of Anopheles beklemishevi and Aedes punctor has been studied. Two new genera of microsporidia are described based on lightmicroscopic and ultrastructural characteristics of spores and sporogony stages. The spore wall of Crepidula beklemishevi gen. n. et sp. n. is formed by two-membrane exospore, thick exospore, bilayer endospore and thin plasmolemma. Spores with single nucleus, polar filament anisofilar, with 6-7 coils (2+ 4-5), polaroplast consisting of three parts: macrochelicoidal, microhelicoidal and lamellar. Fixed spores 4.2 +/- 0.22 x 2 +/- 0.01 microns. The sporogony of Dimeiospora palustris gen. et. n. results in spore formation of two different types. Spores of the first type are oviform, with thick wall, single-nuclear, 6.1 x 4.9 microns. Spore wall with three layers, about 370 nm. Exospore electron-dense, subexospore moderately electrondense. Exospore and subexospore irregularly pleated on the almost spore surface and slightly thinner on anterior end only. Endospore electron-translucent. Polar filament anisofilar, with 9 coils (3 + 6). Polaroplas consists of three parts: lamellar, fine bubbled, and coarse bubbled. Spores of the second type broad-ovate, with apical pole narrower, distal pole concave, 4.6 x 3.7 microns. Spore wall with three layer, 355 nm. Exospore on the apical end irregularly pleated, consists of thin electrondense exospore, subexospore of variable electron density, endospore electron-translucent. Polar filament anisofilar, with 13 coils (3 + 10). Polaroplast has two parts: lamellar and vesicular.     

Epiphytism and terrestrialization in tropicalHuperzia (Lycopodiaceae)

A gap in the exospore and the presence of a mesospore is found to be a normal part of development in the ten species ofSelaginella we have studied. The early spore wall consists of an exospore and a mesospore forming successively on the plasma membrane of the megaspore protoplast when it is 10–15 µm in diameter. Enlargement of the exospore and mesospore creates a central space, the lumen of the megaspore, around the megaspore protoplast. After that there is a vast enlargement of the exospore and a relatively small enlargement of the mesospore. The exospore splits close to its contact with the mesospore forming a gap over equatorial and distal regions. The gap becomes greatly expanded and becomes filled with lipids, PAS-positive carbohydrates, proteins and is crossed by wicks. Experiments with solutions of different osmolality on fresh megaspores show that the exospore and mesospore are not osmotic barriers. The mesospore appears not to be resistant to acetolysis at the many stages tested but exospore is resistant. Thus the mesospore size and shape is retained by the inner exospore that enveloped the mesospore. At maturation the mesospore undergoes lysis and absorption. At the beginning of germination stages an endospore forms at the inner part of the exospore. This inner part of the exospore, that adhered to and enveloped the mesospore, becomes pressed near to the bulk of the exospore. Until pregermination stages the megaspore protoplast is small (10–20 µm in diameter).